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Year-round shellfish exploitation in the Levant and implications for Upper Palaeolithic hunter-gatherer subsistence Authors Marjolein D. Boscha,b*, Marcello A. Manninoc,a, Amy L. Prendergastd, Frank P. Wesselinghe, Tamsin C. O’Connellf and Jean-Jacques Hublina a Department of Human Evolution, Max Planck Institute for Evolutionary Anthropology, Deutscher Platz 6, D-04103 Leipzig, Germany b McDonald Institute for Archaeological Research, University of Cambridge, DoWning Street, Cambridge, CB2 3ER, United Kingdom c Department of Archaeology and Heritage Studies, School of Culture and Society, Aarhus University, Moesgård Allé 20, DK-8270 Højbjerg, Denmark d School of Geography, The University of Melbourne, 221 Bouverie St, Carlton, 3053, Australia e Naturalis Biodiversity Center, Postbus 9517, 2300 RA Leiden, the Netherlands f Department of Archaeology and Anthropology, University of Cambridge, Downing Street, Cambridge CB2 3DZ, United Kingdom *corresponding author: [email protected] Keywords Subsistence behavior, Levantine Upper Paleolithic, shellfish exploitation, oxygen isotope analysis, seasonality, Ksâr ‘Akil, Phorcus turbinatus Highlights • Oxygen isotope analysis conveys year-round shellfish exploitation at Ksâr ‘Akil • Coastal resources had a central role in Early Upper Paleolithic foraging strategies • During most of the Upper Paleolithic, Ksâr ‘Akil was inhabited throughout the year • The EUP occupants of Ksâr ‘Akil had a broader diet than their IUP forbearers Abstract Recent studies have shown that the use of aquatic resources has greater antiquity in hominin diets than previously thought. At present, it is unclear when hominins started to habitually consume marine resources. This study examines shellfish exploitation from a behavioural ecology perspective, addressing how and When past hunter-gatherers from the Levant used coastal resources for subsistence purposes. We investigate the seasonality of shellfish exploitation in the Levantine Upper Palaeolithic through oxygen isotope analysis on shells of the intertidal rocky shore mollusc Phorcus (Osilinus) turbinatus from the key site Ksâr ‘Akil (Lebanon). At this rockshelter, multi-layered archaeological deposits contained remains of both marine and terrestrial molluscs in relatively large quantities, which were consumed and used as tools and ornaments by the occupants of the site. Our results indicate that at the start of the Initial Upper Palaeolithic (IUP), there is no evidence for shellfish consumption. Humans started to take fresh shellfish to the rockshelter from the second half of the IUP onward, albeit in low quantities. During the Early Upper Palaeolithic (EUP) shellfish exploitation became increasingly frequent. Oxygen isotope data show that shellfish exploitation Was practised in every season throughout most of the Upper Palaeolithic (UP), with an emphasis on the colder months. This suggests that coastal resources had a central role in early UP foraging strategies, rather than a seasonally restricted supplementary one. Year- round shellfish gathering, in turn, suggests that humans occupied the rockshelter at different times of the year, although not necessarily continuously. Our oxygen isotope data is complemented with broader-scale exploitation patterns of faunal resources, both vertebrate and invertebrate, at the site. The inclusion of coastal marine resources signifies a diversification of the human diet from the EUP onward, Which is also observed in foraging practices linked to the exploitation of terrestrial fauna. 1 Introduction Recent archaeological discoveries indicate that the exploitation of aquatic resources has a long history among various hominin species. The earliest evidence of the exploitation of freshWater fish was documented from Koobi Fora, Kenya about 1.95 million years ago (Braun et al. 2010) for Homo ergaster/erectus, who made OldoWan tools. Homo erectus individuals may have consumed freshWater molluscs from approximately ≥ 400 thousand years ago (kya) in Trinil (Java, Indonesia) (Joordens et al. 2014). During Marine Isotope Stages (MIS) 6 and 5 (roughly 190–125 kya and 125–80 kya, respectively), Middle Stone Age (MSA) Homo sapiens exploited shellfish at several South African coastal sites (e.g. Klein et al. 2004; Avery et al. 2008; Jerardino and Marean 2010; Langejans et al. 2012; Clark and Kandel 2013; Kyriacou et al. 2015; Jerardino 2016). In the Mediterranean region, marine molluscs were exploited during the Middle Palaeolithic both by H. sapiens and Homo neanderthalensis from MIS 5 onward (e.g., Emiliani et al. 1964; Klein and Scott 1986; Stiner 1999; Finlayson et al. 2006, 2008; Colonese et al. 2011; Cortés-Sánchez et al. 2011; Barker et al. 2012; Fa et al. 2016). Thus, aquatic resources played a role in early hominin diets, although their contribution has been considered as marginal for most of the Palaeolithic (e.g., Erlandson and Moss 2001; Colonese et al. 2011; Clark and Kandel 2013; Jerardino 2015). This is especially true when optimal foraging models are used, which primarily consider the caloric value and energy intake of foodstuffs (see also Grayson and Delpech 1998; Stiner, 2001, 2010). HoWever, from a nutritional ecology perspective, shellfish are a rich source of many essential nutrients, including vitamins (i.e., A, B12, C, D, and E), iron, folate, potassium, calcium, and omega-3 fatty acids (e.g., Hockett and HaWs 2003; Haws and Hockett 2004; Cunnane and CraWford 2014; Kyriacou et al. 2015). Due to their high polyunsaturated (or omega-3) fatty acid content in the form of docosahexaenoic acid and its importance for brain development, shellfish and aquatic food sources in general are thought to have been important in human brain expansion and evolution (e.g., Brenna and Carlson 2014; Cunnane and CraWford 2014; Joordens et al. 2014; Kyriacou et al. 2014). With regard to nutritional ecology, Hockett and HaWs (2003; HaWs and Hockett 2004) suggest that broader and nutritionally rich diets increase hominid fitness, aid to reduce child mortality, and inter-birth intervals. This in turn, would allow for population increase as for example the increase in population density thought to coincide With the start of the UP (e.g., Mellars and French 2011; French 2015). Most of the above-mentioned essential nutrients can also be obtained in reasonable quantities through the consumption of animal organs and/or plant foods, with the possible exceptions of vitamins D and omega-3 fatty acids, which are more difficult to obtain from other, terrestrial, sources. HoWever, due to their limited visibility in the archaeological record, it is often hard to assess the role that organ meat and plant foods played in hominin diets. Whereas shellfish are convenient ‘food packages’, providing an array of essential nutrients not combined in other food sources and, if available, are ideal to assess the nutritional breadth of the diet. Here we use oxygen isotope analysis as a tool to determine the nature and seasonal timing of UP shellfish exploitation. Seasonality of shellfish exploitation data, derived from oxygen isotope analysis, alloW us to explore questions about Whether coastal environments Were used as a supplementary resource in lean times, or if they occupied a more central-place in past foraging strategies and accessed throughout the year. We further investigate Whether shellfish gathering Was a result of intensification of food resources With increased residency and or population pressure (e.g., Stiner et al. 2013; Marean 2014) through comparisons with exploitation patterns of other faunal resources. Moreover, data on the periodicity of mollusc collection are helpful for gaining a fuller understanding of coastal foraging strategies (e.g., Mannino et al. 2007, 2014; Colonese et al. 2009; Prendergast et al. 2016) and allow us to address questions regarding behavioural ecology. For example, how prehistoric hunter-gatherers moved in the landscape or how they used their surroundings for raW material procurement (e.g., bivalves of Glycymeris sp. for the manufacture of utilitarian objects or perforated shells for personal ornamentation) or foraging. In addition, these data can be used as a proxy for the timing of site occupation (e.g., Shackleton 1973; Mannino et al. 2011). Here We investigate seasonality of shellfish exploitation by modern humans in the eastern Mediterranean UP using the Ksâr ‘Akil (Lebanon) shell assemblage as a case study. To do this We conducted oxygen isotope analysis of the most common edible marine mollusc, Phorcus turbinatus. These data on coastal foraging are complemented With broader patterns of faunal resource and habitat exploitation. 2 Ksâr ‘Akil and its palaeoenvironmental setting The Ksâr ‘Akil rockshelter is situated close to the eastern Mediterranean coast, approximately 10 km north of Beirut, Lebanon (Fig. 1). Its multilayered sequence is 23 m deep and spans the Middle Palaeolithic (MP) to the Epipalaeolithic (EPI) (e.g., EWing 1947; Mellars and Tixier 1989). It yielded several human fossils (EWing 1960; Bergman and Stinger 1989; Tillier and Tixier 1991) that are associated With rich archaeological assemblages (Fig. 2). Virtually all mollusc remains Were found in UP deposits (van Regteren Altena 1962; Inizan and Gaillard 1978). The UP sequence at Ksâr ‘Akil comprises roughly the upper 16 m of sediments. In this study, the sequence is subdivided folloWing the division in archaeological layers by Williams and Bergman (2010; see also Tixier and Inizan 1981; Azoury 1986; Marks and Volkman 1986; Bergman and Goring-Morris

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